Archivos de Zootecnia ISSN: 0004-0592 [email protected] Universidad de Córdoba España
Carrara, E.R.; Gaya, L.G.; Mourão, G.B. Fatty acid profile in bovine milk: Its role in human health and modification by selection Archivos de Zootecnia, vol. 66, núm. 253, 2017, pp. 151-158 Universidad de Córdoba Córdoba, España
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Fatty acid profile in bovine milk: Its role in human health and modification by selection
Carrara, E.R.1@; Gaya, L.G.1 and Mourão, G.B.2
1Federal University of São João del-Rei. São João del-Rei. Minas Gerais. Brazil. 2“Luiz de Queiroz” College of Agriculture. University of São Paulo. Piracicaba. São Paulo. Brazil.
SUMMARY
Additional keywords Milk fat is predominantly composed of fatty acids. Bovine milk shows a high concentration Dairy cattle. of saturated fatty acids, some of them associated with an increase in the risk of cardiovascular diseases in human beings. However, many of these have neutral or positive effects on health, as Fat milk. well as some unsaturated fatty acids do. The fat profile of bovine milk can be changed by lipid Genetic parameters. supplementation in a fast and efficient way, but genetic variation is also verified in this profile and genetic breeding becomes more relevant than supplementation for presenting permanent results. This bibliographic review has the objective of gathering the main results referring to the genetic aspects of fatty acids profile in bovine milk.
Perfil de ácidos graxos no leite bovino: seu papel na saúde humana e sua modificação por seleção
Palavras-chave adicionais RESUMO
Gado de leite. A gordura do leite é composta, predominantemente, por ácidos graxos. O leite bovino Gordura do leite. apresenta alta concentração de ácidos graxos saturados, alguns associados ao aumento Parâmetros genéticos. do risco de doenças cardiovasculares em humanos. Porém, muitos desses, possuem efeitos neutros ou positivos sobre a saúde, assim como alguns ácidos graxos insaturados. O perfil da gordura do leite de bovinos pode ser alterado por suplementação lipídica de maneira rápida e eficiente, porém, verifica-se também variação genética nesse perfil e o melhoramento gené- tico se torna mais relevante do que a suplementação por apresentar resultados permanentes. Esta revisão de literatura teve como objetivo reunir os principais resultados referentes aos aspectos genéticos do perfil de ácidos graxos no leite de bovinos. Información
Cronología del artículo. Recibido/Received: 12.02.2016 Aceptado/Accepted: 09.09.2016 On-line: 15.01.2017 Correspondencia a los autores/Contact e-mail: [email protected]
INTRODUCTION lipid supplementation (Santos et al., 2001). This altera- tion can be induced by feed because of the amount of The major part of fat in bovine milk is composed unsaturated fatty acids contained in the provided lipid by triacylglycerols, consisting of three fatty acids and and by its release rate in rumen; therefore, modifying one glycerol molecule. Such fatty acids are predomi- the metabolism, it is possible to obtain a different fatty nantly in saturated form (Eifert, 2006), which are as- acids profile. The toxicity of unsaturated fatty acids sociated with the increase of the risk of cardiovas- through acting upon some bacteria species, leads to cular diseases in human beings (Santos et al., 2013). modification of ruminal microbiota. The use of animal However, such effects are only related to lauric (C12:0), breeding stands out among attempts of modifications myristic (C14:0), and palmitic (C16:0) fatty acid, while of this profile, since there is a genetic variation in fat the others have neutral or positive effects on human milk composition and between bovine breeds (Soyeurt health (Mensink et al., 2003). Regarding the unsatura- et al., 2006). Furthermore, even diet supplementation ted fatty acids, those which stand out are the oleic acid being the fastest and most efficient way to modify fatty (C18:1 cis-9) and the isomers of conjugated linoleic acid acids composition in cow milk (Jenkins & McGuire, (CLA), related to the cholesterol reduction and to the 2006), animal breeding becomes more relevant becau- anticarcinogenic effects respectively (Haug et al., 2007). se it shows permanent advances, despite its slowness The fat profile of bovine milk can be altered by dietary and long term planning. Moreover, recent researches
Arch. Zootec. 66 (253): 151-158. 2017 CARRARA, GAYA AND MOURÃO by Bastin et al. (2013) and Bilal et al. (2014) have shown There are reports that the lauric fatty acid (C12:0) can medium and high heritability for several fatty acids in have antibacterial effects (Sun et al., 2002) and might Holstein cow milk, which suggests a feasible selection act as anticaries and antiplaque agent (Sun et al., 2003). to those components. Thus, studies related to genetic The increase in cardiovascular diseases among human behavior of fatty acids profile in milk might allow the beings is related to lauric (C12:0), myristic (C14:0), and use of tools aiming to improve such profile and promo- palmitic fatty acids, because of low density lipopro- te better milk quality. tein (LDL) and high density lipoprotein (HDL), both of them linked to increased cholesterol. Other milk- Therefore, the objective of this revision was to saturated fatty acids have neutral or positive effect on gather the main results regarding the influence of ani- health (Mensink et al., 2003). Stearic fatty acid (C18:0) mal genetics on fatty acids profile of bovine milk. is not regarded as a promoter of increased cholesterol Chemical composition of bovine milk concentration (Grundy, 1994). Bovine milk is composed of water (87%), lactose In relation to unsaturated fatty acids, the trans form (4.9%), fat (3.5%), protein (3.1%), minerals and vitam- shows worse effect in human health than the satura- ins in smaller proportions (Jensen, 1995). It is necessary ted ones, since it increases the levels of what is called to consider milk composition because it is important to “bad” cholesterol (LDL) and it decreases the levels of determine its nutritional quality, capacity of being pro- “good” cholesterol, HDL. However, only small amou- cessed and consumption by humans. It varies influen- nts of natural trans fats are found in meat and ruminant ced by multiple factors (Silva, 1997). Around 60% of the milk, and they are not associated with cardiovascular variation in milk composition is due to genetic aspects diseases (Mozaffarian et al., 2009). The C16:1 trans-7, a and environmental aspects such as nutrition, climate, trans fatty acid that performs as a biomarker of milk diseases, and milk storage (Kitchen, 1981; Shearer et al., fat, relates itself to a lower risk of diabetes mellitus 1992) causing rest. development and sudden cardiac death (Mozaffarian, 2015b). Among bovine milk components, fat presents hig- her variation rates from 3.2% to 6%. About 98% of cow It is important to highlight that there is great dis- milk lipid fraction is composed by triacylglycerol. Ac- tinction between trans fatty acids that are formed by cording to description by Hayes and Khosla (1992), the animals and those industrially produced. Products combination of lipids in milk that is more favorable to originated from ruminants show a concentration from human health stands for 30% of saturated fatty acids, 3 to 8% of trans fatty acids in relation to total fatty 60% of monounsatured fatty, and 10% of polyunsatu- acids, while industrialized products show 10 to 40% rated fatty acids. The highest concentration of bovine concentration (Martinez, 2009). High levels of indus- milk is fatty acids (Eifert, 2006), but there are only trially produced trans fatty acids can be consumed three of them associated with increase in the risk of through partially hydrogenated vegetable oil, which cardiovascular diseases in human beings: lauric acid; usually contain from 30 to 60% of trans fatty acids myristic acid; and palmitic acid (Mensink et al., 2013). (Mozaffarian, 2016). Besides a higher concentration of Researches indicate that, in the milk fat profile, there trans fatty acids in the total fatty acids, industrializa- are compounds that are beneficial to human health tion also causes the preferential production of elaidic (Eifert, 2006), such as some saturated and unsaturated acid (C18:1 trans-9). Rumen bacteria, mainly generate fatty acids, as described below. vaccenic acid (C18:1 trans-11), a precursor of a specific type of conjugated linoleic acid (CLA) (Martinez, 2009). Fatty acids composition and human health The ingestion of trans fatty acids from partially hydro- Researches highlight the increase in risk of cardio- genated oils is consistently associated with the risk of vascular diseases (Grimsgaard et al., 1999; Zheng et cardiovascular diseases and sudden death (Mozaffa- al., 1999; Yli-Jama et al., 2002; Matsumoto et al., 2013) rian et al., 2006; Mente et al., 2009). when there is fatty acid ingestion, but not all studies There are unsaturated fats that have positive effects present the same ideas (Lemaitre et al., 2010; Wu et al., on human health. Polyunsaturated ω-3 fatty acids can 2011). Saturated fat covers compounds that vary from contribute to the treatment or prevention of cancer, ar- 6 to 24 carbons, which originate in several foods and thritis, depression, and Alzheimer’s disease. Polyunsa- have different metabolic performances (Mozaffarian, turated ω-6 fatty acids derivatives of linoleic acid take 2015a). For instance, the palmitic acid (C16:0) might part in cell membrane structures, playing an important show adverse metabolic effects in vitro, while medium physiological role in the human body and influencing chain (C6:0 to C12:0), odd chain (C15:0 to C17:0), and in blood viscosity, as well as vessel permeability, and very long chain (C20:0 to C24:0) saturated fatty acids also in blood pressure, inflammatory reactions, and might, in turn, show metabolic benefits (Forouhiet platelet functions (Moraes & Colla, 2006). al., 2014). Thus, it is incorrect to group all saturated fatty acids based on only one chemical trait, which Special attention has been given to CLA, a group of is the absence of double bonds in the carbon chain isormers of linoleic acid with some of its unsaturations (Mozaffarian, 2015a). The butyric acid (C4:0) can have being combined, in other words, without methylene anticarcinogenic effect (German, 1999), caprylic (C8:0) between them (Roche et al., 2001). Several beneficial and capric (C10:0) acids might play antiviral roles, and effects of the CLA to human health have been reported the caprylic acid can even have functions related to in the last years, such as antidiabetic and anticarci- the delay of tumor growth (Thormar et al., 1994) and nogenic effects. Among other benefits, there are also act against Gram-negative coliform (Sun et al., 2002). modulations of the immune system, energy partition,
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Table I. Heritability values for groups of fatty acids in bovine milk (Valores de herdabilidade para grupos de ácidos graxos no leite bovino). Common denomination of fatty acid Designation Heritability Source Soyeurt et al. (2007) Saturated AGS 0.14 - 0.46 Bastin et al. (2013) Bilal et al. (2014) Soyeurt et al. (2007) Monounsaturated AGMI 0.21 - 0.26 Bastin et al. (2013) Bilal et al. (2014) Bastin et al. (2013) Polyunsaturated AGPI 0.15 - 0.31 Bilal et al. (2014) Unsaturated AGI 0.27 Bastin et al. (2013) and reduction in the development of atherosclerosis the goals of the programs are being expanded to inclu- (Bauman et al., 1999; Zacarchenco et al., 2013). For this de cow’s longevity, continuing production, and, more reason, CLA has become the focus of many researches. recently, quality traits of milk, in relation to human The major sources of CLA in human diet are the milk health, food safety, and industrial yield of dairy pro- and its derived products, and the enrichment of those ducts. foods with CLA can be a very interesting alternative due to its benefits (De Luca & Jenkins, 2000). The most Genetic parameters of fatty acids profile in bovine milk abundant CLA isomer in milk fat is the C18:2 cis-9 trans 11 (rumenic acid), which represents from 75 to 90% of Heritability the total CLA (Bauman et al., 2003). Results show that heritability values have varied Animal breeding is gradually allowing the increase from 0.15 to 0.46 to groups of saturated, monounsa- of milk production. However, due to market changes, turated, polyunsaturated, and unsaturated fatty acids
Table II. Heritability values for diverse saturated fatty acids in bovine milk (Valores de herdabilidade para diversos ácidos graxos saturados no leite bovino). Common denomination of fatty acid Designation Heritability Source Stoop et al. (2008) Butyric acid C4:0 0.08 - 0.37 Bastin et al. (2013) Bilal et al. (2014) Stoop et al. (2008) Caproic acid C6:0 0.14 - 0.46 Bastin et al. (2013) Bilal et al. (2014) Stoop et al. (2008) Caprylic acid C8:0 0.24 - 0.48 Bastin et al. (2013) Bilal et al. (2014) Stoop et al. (2008) Capric acid C10:0 0.34 - 0.54 Bastin et al. (2013) Bilal et al. (2014) Soyeurt et al. (2007) Stoop et al. (2008) Lauric acid C12:0 0.09 - 0.46 Bastin et al. (2013) Bilal et al. (2014) Soyeurt et al. (2007) Stoop et al. (2008) Myristic acid C14:0 0.07 - 0.49 Bastin et al. (2013) Bilal et al. (2014) Soyeurt et al. (2007) Stoop et al. (2008) Palmitic acid C16:0 0.03 - 0.43 Mele et al. (2009) Bastin et al. (2013) Bilal et al. (2014) Bastin et al. (2013) Margaric acid C17:0 0.21 - 0.40 Bilal et al. (2014) Soyeurt et al. (2007) Stoop et al. (2008) Stearic acid C18:0 0.08 - 0.28 Mele et al. (2009) Bastin et al. (2013) Bilal et al. (2014)
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Table III. Heritability values for diverse unsaturated fatty acids in bovine milk (Valores de herdabilidade para diver- sos ácidos graxos insaturados no leite bovino). Common denomination of fatty acid Designation Heritability Source Myristelaidic acid C14:1 trans-9 0.02 Bilal et al. (2014) Mele et al. (2009) Myristoleic acid C14:1 cis-9 0.19 - 0.39 Bilal et al. (2014) Trans-palmitoleic acid C16:1 trans-9 0.05 Bilal et al. (2014) Mele et al. (2009) Palmitoleic acid C16:1 cis-9 0.14 - 0.30 Bilal et al. (2014) Stoop et al. (2008) Mele et al. (2009) Oleic acid C18:1 cis-9 0.17 - 0.22 Bastin et al. (2013) Bilal et al. (2014) Stoop et al. (2008) Elaidic acid C18:1 trans-9 0.01 - 0.11 Bilal et al. (2014) Stoop et al. (2008) Vaccenic acid C18:1 trans-11 0.03 - 0.12 Mele et al. (2009) Bilal et al. (2014) Stoop et al. (2008) Linolelaidic acid C18:2 trans-9,12 0.02 - 0.13 Bilal et al. (2014) Soyeurt et al. (2007) Linoleic acid C18:2 cis-9,12 0.15 Bilal et al. (2014) Stoop et al. (2008) Rumenic acid C18:2 cis-9, trans-11 0.07 - 0.21 Mele et al. (2009) Bilal et al. (2014) Trans-10, cis-12 linoleic acid C18:2 trans-10, cis-12 0.01 Bilal et al. (2014) Stoop et al. (2008) Alpha-linolenic C18:3 cis-9,12,15 0.06 - 0.09 Bilal et al. (2014)
(table I). According to several authors, estimated he- additive genetic effect on these traits, and they can ritability for specific saturated fatty acids have varied respond well to selection. Thus, genetics can be used from 0.03 to 0.54 (table II), and for specific unsaturated in the manipulation of several of the levels of fatty fatty acids, from 0.01 to 0.39 (table III). acids of milk.
The estimates of heritability to various fatty acids Genetic correlations have shown wide variation according to the sources The values for genetic correlation between the fatty researched. This variation occurs most likely because acids groups and the milk production reported by heritability is not a fixed parameter. In other words, researchers vary from -0.48 to 0.15, and between the it can vary according to population diversity, sample fatty acids groups and the fat percentage, from -0.89 size, number and types of environments, precision in to 0.97 (table IV). The specific genetic correlations conducting the experiment and data recording, as well between saturated fatty acids and milk production as estimation of the methods used. vary from -0.50 to 0.30, and those between acids and So far, the researches conduced showed medium fat percentage vary from -0.43 to 0.83 (table V). The and high heritability estimates for many fatty acids genetic correlations between unsaturated fatty acids found in bovine milk, this suggests a larger share of and milk production vary from -0.31 to 0.77, and those
Table IV. Genetic correlation values among fatty acids, milk production, and fat percentage in bovine milk (Valores de correlação genética entre ácidos graxos, produção de leite e porcentagem de gordura no leite bovino). Common denomination of Genetic correlation with Genetic correlation with fat Designation Source fatty acid milk production percentage Soyeurt et al. (2007) Saturated AGS -0.48 to -0.16 0.91 to 0.97 Bastin et al. (2013) Bilal et al. (2014) Soyeurt et al. (2007) Monounsaturated AGMI -0.35 to 0.15 -0.89 to 0.74 Bastin et al. (2013) Bilal et al. (2014) Bastin et al. (2013) Polyunsaturated AGPI -0.37 to 0.19 -0.80 to 0.69 Bilal et al. (2014) Unsaturated AGI -0.36 0.73 Bastin et al. (2013)
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Table V. Genetic correlation values among saturated fatty acids, milk production, and fat percentage in bo- vine milk (Valores de correlação genética entre ácidos graxos saturados, produção de leite e porcentagem de gordura no leite bovino). Genetic correlation with milk Genetic correlation with fat Common denomination of fatty acid Designation Source production percentage Stoop et al. (2008) Butyric acid C4:0 0.05 and 0.09 0.03 and 0.16 Bilal et al. (2014) Stoop et al. (2008) Caproic acid C6:0 -0.28 and 0.01 0.46 and 0.83 Bastin et al. (2013) Stoop et al. (2008) Caprylic acid C8:0 -0.31 and 0.03 0.34 and 0.81 Bastin et al. (2013) Stoop et al. (2008) Capric acid C10:0 -0.32 and 0.10 0.09 and 0.77 Bastin et al. (2013) Bastin et al. (2013) Lauric acid C12:0 -0.34 and -0.26 0.43 and 0.77 Bilal et al. (2014) Stoop et al. (2008) Mele et al. (2009) Myristic acid C14:0 -0.34 and 0.30 -0.43 and 0.04 Bastin et al. (2013) Bilal et al. (2014) Stoop et al. (2008) Palmitic acid C16:0 -0.50 and -0.33 0.65 and 0.74 Mele et al. (2009) Mele et al. (2009) Stearic acid C18:0 -0.30 and -0.20 -0.10 and 0.71 Bastin et al. (2013) Bilal et al. (2014) between acids and fat percentage vary from -0.85 to C14:0 and saturated fatty acids the genetic correlations 0.69 (table VI). were negative. The same occurred with C4:0, C16:0, and C18:0 when genetically correlated to other studied Besides the estimated genetic correlations among fatty acids. fatty acids, milk production, and fat percentage, esti- mates between fatty acids in one to one basis were also Bastin et al. (2013) estimated genetic correlations of related. Stoop et al. (2008) have estimated positive and milk fatty acid among the three first deliveries, finding negative genetic correlations among several fatty acids values from 0.51 to 0.76 between the first and second found in Holstein cow milk and found values ranging birth, from 0.48 to 0.74 between second and third bir- from -0.84 to 0.99, with negative correlations from -0.84 th, and from 0.44 to 0.70 between first and third birth. to -0.01 and positive correlations from 0.01 to 0.99. Po- The results show that the level of fatty acids in milk sitive and strong correlations were reported from C6:0 tend not to vary much among the three first lactations to 14:0 and unsaturated fatty acids, but between C6:0 to so that if a cow shows high levels of fatty acid in the
Table VI. Genetic correlation values among unsaturated fatty acids, milk production, and fat percentage in bovine milk (Valores de correlação genética entre ácidos graxos insaturados, produção de leite e porcentagem de gordura no leite bovino). Common denomination of Genetic correlation with Genetic correlation with fat percent- Designation Source fatty acid milk production age Mele et al. (2009) Myristoleic acid C14:1 cis-9 0.05 -0.05 and 0.10 Bilal et al. (2014) Mele et al. (2009) Palmitoleic acid C16:1 cis-9 0.09 0.24 and 0.34 Bilal et al. (2014) Stoop et al. (2008) Oleic acid C18:1 cis-9 -0.31 to 0.32 -0.85 to 0.64 Bastin et al. (2013) Bilal et al. (2014) Stoop et al. (2008) C18:1 Vaccenic acid -0.07 and 0.34 -0.69 to -0.43 Mele et al. (2009) trans-11 Bilal et al. (2014) C18:2 cis- Stoop et al. (2008) Linoleic acid 0.24 and 0.77 -0.70 and -0.69 9,12 Bilal et al. (2014) Stoop et al. (2008) C18:2 cis-9, Rumenic acid 0.33 to 0.35 -0.68 to -0.55 Mele et al. (2009) trans-11 Bilal et al. (2014) C18:3 cis- Stoop et al. (2008) Alpha-linolenic 0.15 and 0.53 -0.75 and -0.55 9,12,15 Bilal et al. (2014)
Archivos de zootecnia vol. 66, núm. 253, p. 155. CARRARA, GAYA AND MOURÃO first lactation, it will also show high levels in second level value. On the other hand, Khatib et al. (2008) and third lactations. The estimates among fatty acids identified a polymorphism related to lower fat levels varied from 0.31 to 0.99, suggesting that the selection in the replacement of a cytosine (C allele) by a guanine of one fatty acid can affect the others. (G allele) in exon 8 of the same gene, since the G allele is related to this lower level. According to Bilal et al. (2014), genetic correlations estimates among studied fatty acids were from -0.72 to In the bovine chromosome 19 (BTA19), a significant 0.97, with negative correlations from -0.72 to -0.01, and QTL was recognized in the same spot that contains the positive ones from 0.01 to 0.97. Null genetic correlation synthase fatty acid gene (FASN) (Morris et al., 2007). was reported between C18:3 cis-9,12,15 and C12:0 fatty The FASN is an enzyme that catalyzes once again the acids. synthesis of fatty acids in mammals (Ciecierska et al., Like the heritability estimates, there was a variation 2013). For this reason, and also because of its function between genetic correlations and fatty acids, and pro- of fatty acids synthesis, FASN gene is a candidate for duction variables when compared to published resear- some traits of milk production (Morris et al., 2007). ches. In the same way, estimates in different works can According to Ciecierska et al. (2013), one of the FASN differ according to the studied population, the sample polymorphisms is the FASN17924A>G, which cha- size, endogamy, and estimation methodologies used. racterizes only one exchange of the threonine amino acids for the alanine. These authors have identified that According to the published studies about genetic there were relations between FASN17924A>G poly- correlations among fatty acids, it is possible to as- morphism and the fat percentage in Holstein cow milk. sure that a selection directed to one acid affects the others, since it required the precise definition of goals The main transcription factor that regulates the ex- in breeding programs so that there will not be losses pression of genes involved in fatty acids synthesis and in milk quality. A favorable change aiming good fatty cholesterol is the Sterol Regulatory Element Binding acids profile in milk is still uncertain and the objectives Protein (SREBPs; Brown & Goldstein, 1999). Nafikov et should be well defined before those traits are included al. (2013) have showed there is a significant association in breeding programs (Bastin et al., 2013). of SREBF1 haplotypes with concentrations of the C12:0 and C14:0 fatty acids, being the H1 haplotype related Tools for genomic selection applied to fatty acids profile in bovine milk to lower concentrations of those acids. The identification of molecular marker for fatty Genomic selection tools confirm that genetics in- acids profile in milk as well as other techniques linked terferes in milk fatty acids profile so that more resear- to molecular genetics, appears as an assisting tool for ches involving biotechnology and traditional breeding understanding genetic control of traits and, conse- methodologies are required in order to understand this quently, as an auxiliary tool to selection programs. profile formation better. With the growth of researches regarding the de- termination of loci that are responsible for fatty acids FINAL CONSIDERATIONS in milk, and the use of molecular markers, a better understanding about genetic control of these traits is Conducted studies based on traditional breeding expected. Research on genomic selection for fatty acids methodologies and genomic selection assure that there profile in milk is also recent. is an influence of the animal genetic on milk fatty acids Taniguchi et al. (2004) have verified a polymor- profile. Therefore, researches look for better understan- phism linked to the Δ9-desaturase enzyme (SCD1) in ding the behavior of fatty acids production in bovines, bovines of the Japanese Black breed. Schennink et al. aiming better milk quality. (2008) have reported that larger proportions of C10:0; In the studies regarding genetic parameters, Hols- C12:0. C14:0. C16:1; and CLA were associated with C tein animals were used, but it is important to include allele of the locus SCD1, while smaller proportions of other breeds. There have been variations in heritability C10:1; C12:1; C14:1; C18:0; and C18:1 were linked to T estimates and genetic correlations among fatty acids allele of the same locus. in the analyzed studies. Besides, there has also been An example of SNP polymorphism is the exchange divergence in literature regarding beneficial and har- of a cytosine (C allele) for a thymine (T allele), which mful effects in human health related to saturated fatty results in the replacement of a valine by the alanine in acids. Thus, further studies should be conducted about position 293 of the protein (A293V) (Taniguchi et al., genetic parameters and genomic selection on milk fatty 2004). According to Mosley & McGuire (2007), SCD1 acids profile so that the most appropriate selection A293V polymorphism contributes in 32% to 51% when strategies can be determined in breeding programs. it explains genetic variance referring to unsaturation levels of C10. C12, C14, and C16, 18% of CLA, and 6% BIBLIOGRAPHY of C18. Bastin, C.; Soyeurt, H. and Gengler, N. 2013. Genetic parameters of A research by Brym et al. (2004) reports the influen- milk production traits and fatty acid contents in milk for Holstein cows ce of STAT5A – transcription factor and protein acti- in parity 1-3. J Anim Breed Genet, 130: 118-127. vator located in chromosome 19 – on the level of milk Bauman, D.E.; Baumgard, L.H.; Corl, B.A. and Griinari, J.M. 1999. fat. A replacement of an adenine (A allele) by a guanine Biosynthesis of conjugated linoleic acid in ruminants. Proc Am Soc (G allele) in intron 9 has been found, in which samples Anim Sci, pp. 1-15.
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